THE WATER-WATER CYCLE IN CHLOROPLASTS: Scavenging of Active Oxygens and Dissipation of Excess Photons
Tóm tắt
▪ Abstract Photoreduction of dioxygen in photosystem I (PSI) of chloroplasts generates superoxide radicals as the primary product. In intact chloroplasts, the superoxide and the hydrogen peroxide produced via the disproportionation of superoxide are so rapidly scavenged at the site of their generation that the active oxygens do not inactivate the PSI complex, the stromal enzymes, or the scavenging system itself. The overall reaction for scavenging of active oxygens is the photoreduction of dioxygen to water via superoxide and hydrogen peroxide in PSI by the electrons derived from water in PSII, and the water-water cycle is proposed for these sequences. An overview is given of the molecular mechanism of the water-water cycle and microcompartmentalization of the enzymes participating in it. Whenever the water-water cycle operates properly for scavenging of active oxygens in chloroplasts, it also effectively dissipates excess excitation energy under environmental stress. The dual functions of the water-water cycle for protection from photoinihibition are discussed.
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Tài liệu tham khảo
Amako K, 1994, Plant Cell Physiol., 35, 497
Asada K. 1981. Biological carboxylation. InOrganic and Bio-organic Chemistry of Carbon Dioxide, ed. S Inoue, N Yamazaki, pp. 185–251. Tokyo: Kodansha
Asada K. 1992. Production and scavenging of active oxygen in chloroplasts. InMolecular Biology of Free Radical Scavenging Systems, ed. JG Scandalios, pp. 173–92. New York: Cold Spring Harbor Lab. Press
Asada K. 1994. Production and action of active oxygen species in photosynthetic tissues. See Ref.49A, pp. 77–104
Asada K. 1996. Radical production and scavenging in the chloroplasts. InPhotosynthesis and the Environments, ed. NR Baker, pp. 123–50. Dordrecht: Kluwer
Asada K. 1997. The role of ascorbate peroxidase and monodehydroascorbate reductase in H2O2scavenging in plants. See Ref.175, pp. 715–35
Asada K, 1984, Plant Cell Physiol., 25, 1169
Asada K, Endo T, Mano J, Miyake C. 1998. Molecular mechanism for relaxation of and protection from light stress. See Ref.174A, pp. 37–52
Asada K, 1993, Plant Cell Physiol., 34, 39
Asada K, 1992, Plant Cell Physiol., 31, 557
Asada K, Takahashi M. 1987. Production and scavenging of active oxygen in photosynthesis. InPhotoinhibition, ed. DJ Kyle, CB Osmond, CJ Arntzen, pp. 227–87. Amsterdam: Elsevier
Chen G-X, 1989, Plant Cell Physiol., 30, 987
Chen G-X, 1992, Plant Cell Physiol., 33, 117
Chen G-X, 1992, Plant Cell Physiol., 33, 109
Foyer CH. 1997. Oxygen metabolism and electron transport in photosynthesis. See Ref.175, pp. 587–621
Foyer CH, 1994, Causes of Photooxidative Stress and Amelioration of Defense Systems in Plants.
Getzoff ED, 1992, Science, 358, 347
Hossain MA, 1984, Plant Cell Physiol., 25, 85
Hossain MA, 1984, Plant Cell Physiol., 25, 1285
Inoue K, 1986, Plant Cell Physiol., 27, 961
Ivanov BN, 1997, Biokhimiya, 62, 1082
Kanematsu K, 1990, Plant Cell Physiol., 31, 99
Larkum AWD, 1998, Photochem. Photobiol. Biol.
Mano J, 1999, Plant Cell Physiol., 40
Mano J, 1995, Plant Cell Physiol., 36, 1589
Mano J, 1999, Plant Cell Physiol., 40
Mi H, 1995, Plant Cell Physiol., 36, 661
Miyake C, 1992, Plant Cell Physiol., 33, 541
Miyake C, 1993, Plant Cell Physiol., 34, 881
Miyake C, 1991, Plant Cell Physiol., 32, 33
Miyake C, Sano S, Asada K. 1996. A new assay of ascorbate peroxidase using the coupled system with monodehydroascorbate radical reductase. InPlant Peroxidases, Biochemistry and Physiology, ed. C Obinger, U Burner, R Ebermann, C Openel, H Greppin, pp. 386–89. Geneva: Univ. Geneva Press
Miyake C, 1995, Plant Cell Physiol., 36, 743
Mullineaux PM, Creissen GP. 1997. Glutathione reductase: regulation and role in oxidative stress. See Ref.175, pp. 667–713
Nakano Y, 1981, Plant Cell Physiol., 22, 867
Ogawa K, 1997, Plant Cell Physiol., 38, S35
Ogawa K, 1995, Plant Cell Physiol., 36, 565
Sano S, 1994, Plant Cell Physiol., 35, 425
Satoh K, 1998, Stress Responses of Photosynthetic Organisms.
Scandalios JG. 1997. Molecular genetics of superoxide dismutases in plants. InOxidative Stress and the Molecular Biology of Antioxidant Defenses, ed. JG Scandalios, pp. 527–68. NY: Cold Spring Harbor Lab. Press
Schreiber U. 1998. Chlorophyll fluorescence: new instruments for special applications. InProc. Int. Congr. Photosyn., 11th. Dordrecht: Kluwer. In press
Schreiber U, Hormann H, Asada K, Neubauer C. 1995. O2-dependent electron flow in intact spinach chloroplasts: properties and possible regulation of the Mehler-ascorbate peroxidase cycle. See Ref.118A, 2:813–18
Takahashi M, 1982, Plant Cell Physiol., 25, 1457
Takahashi Y, 1984, Plant Cell Physiol., 25, 785
Takeba G, Kozaki A. 1998. Photorespiration is an essential mechanism for the protection of C3plants from photooxidation. See Ref.174A, pp. 37–52
Westphal S, 1992, Z. Naturforsch., 47, 1342
Yamasaki H, Heshiki R, Yamasu T, Sakihama Y, Ikehara N. 1995. Physiological significance of the ascorbate regenerating system for the high-light tolerance of chloroplasts. See Ref.118A, 4:291–94
Zium-Hanck U, 1980, Biochim. Biophys. Acta, 591, 166